Effects of Ageratina Adenophora Invasion on the Understory
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Article Effects of Ageratina adenophora Invasion on the Understory Community and Soil Phosphorus Characteristics of Different Forest Types in Southwest China Xiaoni Wu 1,2,3, Changqun Duan 1,3 , Denggao Fu 1,3,*, Peiyuan Peng 1,3, Luoqi Zhao 1,3 and Davey L. Jones 4,5 1 School of Ecology and Environmental Science & Yunnan Key Laboratory for Plateau Mountain Ecology and Restoration of Degraded Environments, Yunnan University, Kunming 650091, China; [email protected] (X.W.); [email protected] (C.D.); [email protected] (P.P.); [email protected] (L.Z.) 2 School of Agriculture and Life Sciences, Kunming University, Kunming 650214, China 3 International Cooperative Center of Plateau Lake Ecological Restoration and Watershed Management of Yunnan, Yunnan University, Kunming 650091, China 4 School of Natural Sciences, Bangor University, Gwynedd LL57 2UW, UK; [email protected] 5 UWA School of Agriculture and Environment, University of Western Australia, Perth, WA 6009, Australia * Correspondence: [email protected]; Tel./Fax: +86-871-6503-3629 Received: 17 June 2020; Accepted: 23 July 2020; Published: 25 July 2020 Abstract: Understanding the influence of invasive species on community composition and ecosystem properties is necessary to maintain ecosystem functions. However, little is known about how understory plant communities and soil nutrients respond to invasion under different land cover types. Here, we investigated the effects of the invasive species Ageratina adenophora on the species and functional diversity of understory communities and on soil phosphorus (P) status in three forest types: CF, coniferous forest; MF, coniferous and broadleaf mixed forest; and EBF, evergreen broadleaf forest. We found that the species and functional diversity indices of the understory community significantly varied by forest type. Among the invaded plots, the greatest decrease in functional diversity (functional richness, functional divergence, and functional dispersion) and biotic homogenization were found in the CF rather than the MF or EBF. In addition, the invasion by A. adenophora significantly increased the soil NaHCO3-extractable inorganic P and organic P in the MF and EBF, respectively, while obviously decreasing the soil maximum P sorption capacity and maximum buffering capacity in the CF. However, the changes in the species and functional attributes of the understory communities were weakly associated with changes in the soil P status, probably because of the different response times to invasion in different forest types. The implication of these changes for ecosystem structure and function must be separately considered when predicting and managing invasion at a landscape scale. Keywords: biological invasion; functional diversity; understory community; soil phosphorus fractions; soil phosphorus sorption characteristics 1. Introduction Alien plant species invasion is recognized as a serious threat to biodiversity and ecosystem functions [1–3]. Invasive plants can affect natural and semi-natural habitats by displacing native species and changing the nutrient status of the soil [4,5]. Some studies have reported that non-native species invasion may affect terrestrial ecosystem processes and functions via changes in plant community Forests 2020, 11, 806; doi:10.3390/f11080806 www.mdpi.com/journal/forests Forests 2020, 11, 806 2 of 14 composition or soil properties [6,7]. However, many of the observed impact patterns may depend on many confounding factors of the landscape, including the influence of different land cover types [8]. Thus, understanding the influence of invasive species on biodiversity and ecosystem functions in different land cover types can improve spread predictions and reduce ecosystem impacts due to invasive plant species. The effect of invasion is frequently associated with biodiversity loss; however, changes in the plant species composition and diversity may take many years to play out, especially in forest ecosystems. However, increasingly, research is devoted to plant functional traits, exploring how changes in the functional attributes of plant communities affect ecosystem functions and their response to environmental changes [9,10]. Furthermore, trait-based approaches are being used to examine how invasion affects ecosystem structure and functions [11,12]. In forest ecosystems, invasive species may have a detrimental effect on the understory vegetation, which in turn plays a critical role in ecosystem functions [13,14]. Hence, analysing the response of plant functional attributes within the understory community to invasion and the relationships between functional attributes and ecosystem functions might help detect early vegetation responses and ecosystem function alterations. However, the responses of plant functional attributes to invasion and their feedback to ecosystem processes and functions is likely to vary among different forest types. Among ecosystem processes and functions, the effect of invasion on the soil phosphorus (P) cycle is less understood than the effect of invasion on the carbon (C) and nitrogen (N) cycles [15–17]. Most studies on the modification of the P cycle due to exotic species typically focus on the total P (TP) and inorganic P (Pi) forms [18], and less data exist on the impacts of exotic plants on P fractions and P dynamics. Among soil P fractions, Pi extracted with deionized water (water-Pi) and NaHCO3 (bicarb-Pi) are considered the most biologically and readily available Pi forms, and Po extracted with NaHCO3 (bicarb-Po) is easily mineralizable and may contribute to plant-available Pi [18]. In invaded ecosystems, exotic species can affect the distribution and fluxes of easily available P in soil at short- and medium-term time scales, by changing the soil microbial community or soil physicochemical properties [19–21]. However, different patterns of changes in the soil P fraction have been observed in some studies, because the direction and degree of changes in P cycling may depend on many confounding factors, including the specifics of site conditions, land use type, and the biological characteristics of exotic species [22–24]. Therefore, the measurement of easily available P and P behaviour characteristics, together with vegetation properties, is required to better understand the relationships among invasion, vegetation composition, and soil P cycling. The invasive herb Ageratina adenophora, a perennial shrub native to Mexico, invaded China in the 1940s from Burma and is now widespread in Southwest China [22]. In this region, invasion by A. adenophora has a profound influence on the composition, structure, and function of the impacted forest ecosystems because of its clonal reproduction and competitive advantage [16,25]. However, there are few quantitative data on the effects of A. adenophora on understory community composition and soil P status in different forest types. In this study, three typical forest types in Xishan National Forest Park bordering Dianchi Lake, a substantially eutrophic water body in Southwest China, were selected to investigate the invasion effect on understory communities and soil P status, P fraction and P sorption characteristics. Our aims were (1) to determine the effects of invasion by A. adenophora on the understory plant community and soil P status (including the easily available P fractions and P sorption characteristics) in the three forest types, and (2) to quantify the effects of invasion on the relationships between understory plant community properties and soil P status. 2. Materials and Methods 2.1. Site Description The study was carried out at Xishan National Forest Park (102◦37~380 E, 24◦57~590 N), nearby Kunming city, Yunnan Province, China. This park borders Dianchi Lake to the east. Owing to the Forests 2020, 11, 806 3 of 14 influenceForests 2019 of,the 10, x southwestern FOR PEER REVIEW monsoon climate, the average annual precipitation in the area is 11003 of mm.14 The rainy season lasts from May to October each year. The mean annual temperature is 14.7 ◦C. Themm. soils The in rainy the study season area lasts are from classified May to October as Cambisols each year. (according The mean to annual FAO/UNESCO temperature classifications), is 14.7 °C. whichThe developedsoils in the fromstudy basaltarea are parent classified material. as Ca Thembisols original (according vegetation to FAO/UNESCO was a semi-humid classifications), evergreen broadleavedwhich developed forest, from some basalt of which parent was material. utilized The as coppicesoriginal vegetation for fuelwood was a after semi-humid deforestation evergreen before thebroadleaved 1960s. Since forest, the 1980s,some of some which of was these utilized have beenas coppices planted for by fuelwoodPinus armandii after deforestationand P. yunnanensis before afterthedeforestation. 1960s. Since the Due1980s, to some the long-termof these have preservation been planted of by some Pinus originalarmandii vegetationand P. yunnanensis and di afterfferent restorationdeforestation. measures, Due to thethe diversitylong-term ofpreservation vegetation of shows some original a patchy vegetation distribution. and different The main restoration vegetation measures, the diversity of vegetation shows a patchy distribution. The main vegetation types are types are semi-humid evergreen broadleaf forest, coniferous and broadleaf mixed forest, and subtropical semi-humid evergreen broadleaf forest, coniferous and broadleaf mixed forest, and subtropical coniferous forest. In this area, these three forest types with similar conditions,